Problems with comparative analyses of avian brain size

There are multiple hypotheses for the evolution of cognition. The most prominent hypotheses are the Social Intelligence Hypothesis (SIH) and the Ecological Intelligence Hypothesis (EIH), which are often pitted against one another. These hypotheses tend to be tested using broad-scale comparative studies of brain size, where brain size is used as a proxy of cognitive ability, and various social and/or ecological variables are included as predictors. Here, we test how methodologically robust such analyses are. First, we investigate variation in brain and body size measurements across >1000 species of bird. We demonstrate that there is substantial variation in brain and body size estimates across datasets, indicating that conclusions drawn from comparative brain size models are likely to differ depending on the source of the data. Following this, we subset our data to the Corvides infraorder and interrogate how modelling decisions impact results. We show that model results change substantially depending on variable inclusion, source and classification. Indeed, we could have drawn multiple contradictory conclusions about the principal drivers of brain size evolution. These results reflect recent concerns that current methods in comparative brain size studies are not robust. We add our voices to a growing community of researchers suggesting that we move on from using such methods to investigate cognitive evolution. We suggest that a more fruitful way forward is to instead use direct measures of cognitive performance to interrogate why variation in cognition arises within species and between closely related taxa.

Approach direction and accuracy, but not response times, show spatial-numerical association in chicks

Chicks trained to identify a target item in a sagittally-oriented series of identical items show a higher accuracy for the target on the left, rather than that on the right, at test when the series was rotated by 90°. Such bias seems to be due to a right hemispheric dominance in visuospatial tasks. Up to now, the bias was highlighted by looking at accuracy, the measure mostly used in non-human studies to detect spatial numerical association, SNA. In the present study, processing by each hemisphere was assessed by scoring three variables: accuracy, response times and direction of approach. Domestic chicks were tested under monocular vision conditions, as in the avian brain input to each eye is mostly processed by the contralateral hemisphere. Four-day-old chicks learnt to peck at the 4th element in a sagittal series of 10 identical elements. At test, when facing a series oriented fronto-parallel, birds confined their responses to the visible hemifield, with high accuracy for the 4th element. The first element in the series was also highly selected, suggesting an anchoring strategy to start the proto-counting at one end of the series. In the left monocular condition, chicks approached the series starting from the left, and in the right monocular condition, they started from the right. Both hemispheres appear to exploit the same strategy, scanning the series from the most lateral element in the clear hemifield. Remarkably, there was no effect in the response times: equal latency was scored for correct or incorrect and for left vs. right responses. Overall, these data indicate that the measures implying a direction of choice, accuracy and direction of approach, and not velocity, i.e., response times, can highlight SNA in this paradigm. We discuss the relevance of the selected measures to unveil SNA.

Bird neurocranial and body mass evolution across the end-Cretaceous mass extinction: The avian brain shape left other dinosaurs behind

Birds today are the most diverse clade of terrestrial vertebrates, and understanding why extant birds (Aves) alone among dinosaurs survived the Cretaceous-Paleogene mass extinction is crucial to reconstructing the history of life. Hypotheses proposed to explain this pattern demand identification of traits unique to Aves. However, this identification is complicated by a lack of data from non-avian birds. Here, we interrogate survivorship hypotheses using data from a new, nearly complete skull of Late Cretaceous (~70 million years) bird Ichthyornis and reassess shifts in bird body size across the Cretaceous-Paleogene boundary. Ichthyornis exhibited a wulst and segmented palate, previously proposed to have arisen within extant birds. The origin of Aves is marked by larger, reshaped brains indicating selection for relatively large telencephala and eyes but not by uniquely small body size. Sensory system differences, potentially linked to these shifts, may help explain avian survivorship relative to other dinosaurs.

The dorsal arcopallium of chicks displays the expression of orthologs of mammalian fear related serotonin receptor subfamily genes

Fear is an adaptive emotion that elicits defensive behavioural responses against aversive threats in animals. In mammals, serotonin receptors (5-HTRs) have been shown to modulate fear-related neural circuits in the basolateral amygdala complex (BLA). To understand the phylogenetic continuity of the neural basis for fear, it is important to identify the neural circuit that processes fear in other animals. In birds, fear-related behaviours were suggested to be processed in the arcopallium/amygdala complex and modulated by the serotonin (5-HT) system. However, details about the distribution of 5-HTRs in the avian brain are very sparsely reported, and the 5-HTR that is potentially involved in fear-related behaviour has not been elucidated. In this study, we showed that orthologs of mammalian 5-HTR genes that are expressed in the BLA, namely 5-HTR1A, 5-HTR1B, 5-HTR2A, 5-HTR2C, 5-HTR3A, and 5-HTR4, are expressed in a part of the chick arcopallium/amygdala complex called the dorsal arcopallium. This suggests that serotonergic regulation in the dorsal arcopallium may play an important role in regulating fear-related behaviour in birds. Our findings can be used as a basis for comparing the processing of fear and its serotonergic modulation in the mammalian amygdala complex and avian arcopallium/amygdala complex.

As above, so below: Whole transcriptome profiling supports the continuum hypothesis of avian dorsal and ventral pallium organization

Over the last two decades, beginning with the Avian Brain Nomenclature Forum in 2000, major revisions have been made to our understanding of the organization and nomenclature of the avian brain. However, there are still unresolved questions on avian pallial organization, particularly whether the cells above the ventricle represent different populations to those below it. Concerns included limited number of genes profiled, biased selection of genes, and potential independent origins of cell types in different parts of the brain. Here we test two competing hypotheses, using RNA sequencing to profile the transcriptomes of the major avian pallial subdivisions dorsal and ventral to the ventricle boundary, and a new zebra finch genome assembly containing about 22,000 annotated, complete genes. We found that the transcriptomes of neural populations below and above the ventricle were remarkably similar. What had been previously named hyperpallium densocellulare above the ventricle had nearly the same molecular profile as the mesopallium below it; the hyperpallium apicale above was highly similar to the nidopallium below; the primary sensory intercalated hyperpallium apicale above was most similar to the sensory population below, although more divergent than the other populations were to each other. These shared population expression profiles define unique functional specializations in anatomical structure development, synaptic transmission, signaling, and neurogenesis. These findings support the continuum hypothesis of avian brain subdivisions above and below the ventricle space, with the pallium as a whole consisting of four major cell populations instead of seven and has some profound implications for our understanding of vertebrate brain evolution.

The Role of Hp-NCL Network in Goal-Directed Routing Information Encoding of Bird: A Review

Goal-directed navigation is a crucial behavior for the survival of animals, especially for the birds having extraordinary spatial navigation ability. In the studies of the neural mechanism of the goal-directed behavior, especially involving the information encoding mechanism of the route, the hippocampus (Hp) and nidopallium caudalle (NCL) of the avian brain are the famous regions that play important roles. Therefore, they have been widely concerned and a series of studies surrounding them have increased our understandings of the navigation mechanism of birds in recent years. In this paper, we focus on the studies of the information encoding mechanism of the route in the avian goal-directed behavior. We first summarize and introduce the related studies on the role of the Hp and NCL for goal-directed behavior comprehensively. Furthermore, we review the related cooperative interaction studies about the Hp-NCL local network and other relevant brain regions supporting the goal-directed routing information encoding. Finally, we summarize the current situation and prospect the existing important questions in this field. We hope this paper can spark fresh thinking for the following research on routing information encoding mechanism of birds.

Head-direction coding in the hippocampal formation of birds

Birds strongly rely on spatial memory and navigation. However, it is unknown how space is represented in the avian brain. Here we used tetrodes to record neurons from the hippocampal formation (HPF) of Japanese quails – a migratory ground-dwelling species – while the quails roamed a 1×1-meter arena (>2,100 neurons from 21 birds). Whereas spatially-modulated cells (place-cells, border-cells, etc.) were generally not encountered, the firing-rate of 12% of the neurons was unimodally and significantly modulated by the head-azimuth – i.e. these were head-direction cells (HD cells, n=260). Typically, HD cells were maximally active at one preferred-direction and minimally at the opposite null-direction, with preferred-directions spanning all 360°. The HD tuning was relatively broad (mean= 130°), independent of the animal’s position and speed, and was stable during the recording-session. These findings support the existence of an allocentric head-direction representation in the quail HPF, and provide the first demonstration of head-direction cells in birds.

Investigation of stereotactic surgery for avian brain stimulation by a fully implanted wireless system

OBJECTIVEThe authors’ goal was to study avian motor brain mapping via wireless stimulation to induce certain behaviors. In this paper, the authors propose an electrode design that is suitable for avian brain stimulation as well as a stereotactic implant procedure for the proposed electrode.METHODSAn appropriate breed for avian brain study was chosen. A fully implantable remote-controlled electrical stimulation system was inserted to minimize discomfort. A suitable electrode design and stereotactic surgery method based on the electrode design were investigated.RESULTSUsing a wireless stimulation system, flapping and rotation behaviors were induced by stimulating the ventral part of the nucleus intercollicularis and formatio reticularis medialis mesencephali both on the ground and during flight.CONCLUSIONSThe authors were able to implant the entire brain stimulation system inside the avian body without any surgical complications. Postoperative observations suggested that the bird did not find the implant uncomfortable.